Biology:Lamiaceae

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Lamiaceae or Labiatae is a family of flowering plants commonly known as mints, deadnettles, or sages. Many species in Lamiaceae are aromatic, and the family includes many widely used culinary herbs like basil, mint, rosemary, sage, savory, marjoram, oregano, hyssop, thyme, lemon balm, lavender, and perilla, as well as traditional medicines such as catnip, skullcap, bee balm, wild dagga, self-heal, and oriental motherwort.

Some species are shrubs, trees (such as teak), or, rarely, vines, but most are herbaceous. Many members of the family are widely cultivated, not only for their aromatic qualities, but also their ease of cultivation through stem cuttings.[1] Many species are cultivated for their aromatic leaves and can have additional edible parts. Some species are grown for seed, such as Salvia hispanica (chia), or for their edible tubers, such as Plectranthus edulis, P. esculentus, P. rotundifolius, and Stachys affinis (Chinese artichoke). Many are also grown ornamentally, notably coleus, Plectranthus, and Salvia species/hybrids.

The mint family as a whole has a cosmopolitan distribution[2] comprising about 236 genera[3] and around 6,900[2] to 7,200[3] species. The World Checklist of Selected Plant Families estimates an even larger number with 7,534 species.[4] The largest genera are Salvia (900), Scutellaria (360), Stachys (300), Plectranthus (300), Hyptis (280), Teucrium (250), Vitex (250), Thymus (220), and Nepeta (200).[3] Clerodendrum was once a genus of over 400 species,[3] but by 2010, it had been narrowed to about 150.[5]

The family has traditionally been considered closely related to Verbenaceae;[3] in the 1990s, phylogenetic studies suggested that many genera classified in Verbenaceae should be classified in Lamiaceae[6][7] or to other families in the order Lamiales.[8]

The alternative family name Labiatae refers to the flowers typically having petals fused into an upper lip and a lower lip (labia in Latin). Although this is still considered an acceptable alternative name, most botanists now use the name Lamiaceae in referring to this family. The flowers are bilaterally symmetrical with five united petals and five united sepals. They are usually bisexual and verticillastrate (a flower cluster that looks like a whorl of flowers, but actually consists of two crowded clusters). The leaves emerge oppositely, each pair at right angles to the previous one (decussate) or whorled. The stems are frequently square in cross section,[9] but this trait is not found in all members of the family, and is sometimes found in other plant families.

Genera

The last revision of the entire family was published in 2004.[3] It described and provided keys to 236 genera. These are marked with an asterisk (*) in the list below. A few genera have been established or resurrected since 2004. These are marked with a plus sign (+). Other genera have been synonymised. These are marked with a minus sign (-). The remaining genera in the list are mostly of historical interest only and are from a source that includes such genera without explanation.[10] Few of these are recognized in modern treatments of the family.

Kew Gardens provides a list of genera that includes additional information.[11] A list at the Angiosperm Phylogeny Website is frequently updated.[12] Plants of the World Online currently accepts 224 genera.[13]

Recent changes

The circumscription of several genera has changed since 2004. Tsoongia, Paravitex, and Viticipremna have been sunk into synonymy with Vitex.[14] Huxleya has been sunk into Volkameria.[5] Kalaharia, Volkameria, Ovieda, and Tetraclea have been segregated from a formerly polyphyletic Clerodendrum.[5] Rydingia has been separated from Leucas.[15] The remaining Leucas is paraphyletic over four other genera.[16]

Subfamilies and tribes

In 2004, the Lamiaceae were divided into seven subfamilies, plus 10 genera not placed in any of the subfamilies.[3] The unplaced genera were: Tectona, Callicarpa, Hymenopyramis, Petraeovitex, Peronema, Garrettia, Cymaria, Acrymia, Holocheila, and Ombrocharis. The subfamilies are the Symphorematoideae, Viticoideae, Ajugoideae, Prostantheroideae, Nepetoideae, Scutellarioideae, and Lamioideae. The subfamily Viticoideae is probably not monophyletic.[14] The Prostantheroideae, Nepetoideae, Ajugoideae, and Lamioideae are divided into tribes.[17][18] These are shown in the phylogenetic trees below.

By 2026, the APG system has recognized 12 subfamilies; now, Callicarpa is classified in subfam. Callicarpoideae, Acrymia and Cymaria in subfam. Cymarioideae, Holocheila in subfam. Lamioideae, Ombrocharis in subfam. Nepetoideae, Garretia, Hymenopyramis, Peronema and Petraeovitex in subfam. Peronematoideae and Tectona in subfam. Tectonoideae.[19]

Phylogeny

Most of the genera of Lamiaceae have never been sampled for DNA for molecular phylogenetic studies.[18] Most of those that have been are included in the following phylogenetic tree. The phylogeny depicted below is based on seven different sources.[3][7][5][14][20][21][22]

Lamiaceae
 

Callicarpa

 
 

Tectona

 
Viticoideae (pro parte)
 

Gmelina

 
 

Premna

 
 
Viticoideae (pro parte)

Vitex

 
Symphorematoideae
 

Congea

 
 

Symphorema

 
 
Ajugoideae
 

Rotheca

 
 
 

Teucrium

 
 
 

Ajuga

 
 
 
 

Oxera

 
 

Faradaya

 
 
 
 

Kalaharia

 
 
 

Clerodendrum

 
 
 

Volkameria

 
 
 

Ovieda

 
 

Aegiphila

 
 

Tetraclea

 
 

Amasonia

 
 
 
 
 
 
 
 
 
Prostantheroideae
Chloantheae

Chloanthes

 
Westringieae
 

Prostanthera

 
 

Westringia

 
 
 
Nepetoideae
Ocimeae
 

Lavandula

 
 

Siphocranion

 
 
 

Isodon

 
 

Hanceola

 
 

Hyptis

 
 
 
 

Orthosiphon

 
 

Ocimum

 
 
 
 

Plectranthus

 
 

Coleus

 
 
 
 
 
Elsholtzieae
 

Elsholtzia

 
 

Perilla

 
 
Mentheae
 
 

Lepechinia

 
 

Salvia

 
 

Rosmarinus

 
 
 
 
 

Prunella

 
 
 

Nepeta

 
 

Dracocephalum

 
 

Agastache

 
 
 
 
 
 

Origanum

 
 

Thymus

 
 
 
 

Mentha

 
 
 

Satureja

 
 
 

Clinopodium

 
 

Bystropogon

 
 
 
 

Pycnanthemum

 
 

Monarda

 
 
 
 

Dicerandra

 
 

Conradina

 
 
 
 
 
 
 
 
 
Scutellarioideae
 

Holmskioldia

 
 

Scutellaria

 
 
Lamioideae
 

Pogostemon

 
 
 

Phlomis

 
 

Lamium

 
 
 

Stachys

 
 

Sideritis

 
 
 

Haplostachys

 
 
 

Stenogyne

 
 

Phyllostegia

 
 
 
 
 

Leonurus

 
 

Marrubium

 
 

Moluccella

 
 
 

Rydingia

 
 
 

Leucas

 
 

Leonotis

 
 
 
 
 
 
 

Some more recent studies have focused on clarifying the subfamilial relationships within the family using large chloroplast gene datasets with largely congruent results.[17][18]

Li et al. 2016 (dataset D270)[17]
Lamiaceae
 
Subfamily
 

Tribe Westringieae

 
 

Tribe Chloantheae

 
Prostantheroideae
 

Callicarpa

 
 
 
 
 

Subfamily Symphorematoideae

 
 

Subfamily Viticoideae

 
 
Subfamily
 

Tribe Elsholtzieae

 
 
 

Tribe Ocimeae

 
 

Tribe Mentheae

 
 
Nepetoideae
 
 

Tectona

 
 
 

Subfamily Premnoideae

 
 

Subfamily Ajugoideae

 
 
 

Subfamily Peronematoideae

 
 
 

Subfamily Scutellarioideae

 
 
 

Subfamily Cymarioideae

 
Subfamily
 

Tribe Pogostemoneae

 
 
 

Tribe Gomphostemmateae

 
 
 

Tribe Synandreae

 
 
 

Tribe Stachydeae

 
 
 

Tribe Paraphlomideae

 
 
 

Tribe Phlomideae

 
 
 

Tribe Leonureae

 
 
 

Tribe Marrubieae

 
 
 

Tribe Lamieae

 
 

Tribe Leucadeae

 
 
 
 
 
 
 
 
 
Lamioideae
 
 
 
 
 
 
 
Zhao et al. 2021[18]
Lamiaceae
 
Subfamily
 

Tribe Westringieae

 
 

Tribe Chloantheae

 
Prostantheroideae
 

Subfamily Callicarpoideae

 
 
 
Subfamily
 

Tribe Mentheae

 
 
 

Tribe Ocimeae

 
 

Tribe Elsholtzieae

 
 
Nepetoideae
 
 
 

Subfamily Symphorematoideae

 
 

Subfamily Viticoideae

 
 
 
 

Subfamily Tectonoideae

 
 
 

Subfamily Premnoideae

 
 
Subfamily
 

Tribe Rotheceae

 
 
 

Tribe Teucrieae

 
 
 

Tribe Ajugeae

 
 

Tribe Clerodendreae

 
 
 
Ajugoideae
 
 

Subfamily Peronematoideae

 
 
 

Subfamily Scutellarioideae

 
 
 

Subfamily Cymarioideae

 
Subfamily
 

Tribe Pogostemoneae

 
 
 

Tribe Gomphostemmateae

 
 
 

Tribe Colquhounieae

 
 
 

Tribe Synandreae

 
 
 
 

Tribe Stachydeae

 
 
 

Tribe Galeopseae

 
 

Tribe Betoniceae

 
 
 
 
 

Tribe Paraphlomideae

 
 
 

Tribe Phlomideae

 
 
 

Tribe Leonureae

 
 
 

Tribe Marrubieae

 
 
 

Tribe Leucadeae

 
 

Tribe Lamieae

 
 
 
 
 
 
 
 
 
 
Lamioideae
 
 
 
 
 
 
 
 
 

References

  1. ↑ Retief, Elizabeth (July 2008). "Lamiaceae". https://pza.sanbi.org/lamiaceae. 
  2. ↑ 2.0 2.1 Heywood, Vernon H.; Brummitt, Richard K.; Seberg, Ole; Culham, Alastair (2007). Flowering Plant Families of the World. Ontario, Canada: Firefly Books. ISBN 978-1-55407-206-4. 
  3. ↑ 3.0 3.1 3.2 3.3 3.4 3.5 3.6 3.7 Harley RM, Atkins A, Budantsev AL, et al. 2004. "Labiatae" pages 167-275. In: Klaus Kubitzki (editor) and Joachim W. Kadereit (volume editor). The Families and Genera of Vascular Plants volume VII. Springer-Verlag: Berlin; Heidelberg, Germany. ISBN 978-3-540-40593-1
  4. ↑ "World Checklist of Selected Plant Families". https://apps.kew.org/wcsp/incfamilies.do. 
  5. ↑ 5.0 5.1 5.2 5.3 Yuan, Yao-Wu; Mabberley, David J.; Steane, Dorothy A.; Olmstead, Richard G. (2010). "Further disintegration and redefinition of Clerodendrum (Lamiaceae): Implications for the understanding of the evolution of an intriguing breeding strategy". Taxon 59 (1): 125–133. doi:10.1002/tax.591013. Bibcode: 2010Taxon..59..125Y. 
  6. ↑ Cantino, P.D., Harley, R.M. & Wagstaff, S.J. 1992. Genera of Labiatae: status and classification. Pp. 511-522. In: Raymond M. Harley and Tom Reynolds (editors). Advances in Labiate Science. Richmond, Royal Botanic Gardens, Kew.
  7. ↑ 7.0 7.1 Wagstaff, Steven J.; Hickerson, Laura; Spangler, Russ; Reeves, Patrick A.; Olmstead, Richard G. (1998). "Phylogeny in Labiatae s.l., inferred from cpDNA sequences". Plant Systematics and Evolution 209 (3–4): 265–274. doi:10.1007/bf00985232. Bibcode: 1998PSyEv.209..265W. 
  8. ↑ Cite error: Invalid <ref> tag; no text was provided for refs named MOBOT
  9. ↑ Parnell, J. and Curtis, T. 2012. Webb's An Irish Flora. Cork University Press ISBN 978-185918-4783
  10. ↑ "List of genera in Lamiaceae". In: "Lamiaceae". In: "List of families". In: "Families and genera in GRIN. (see External links below)
  11. ↑ List of Genera in Lamiaceae. At: Vascular Plant Families and Genera. At: World Checklist of Selected Plant Families. At: Electronic Plant Information Center. At: Website of Royal Botanic Gardens, Kew. (see External Links below).
  12. ↑ "APG list of Lamiaceae genera". https://www.mobot.org/mobot/research/apweb/genera/lamiaceaegen.html. 
  13. ↑ Cite error: Invalid <ref> tag; no text was provided for refs named powo
  14. ↑ 14.0 14.1 14.2 Bramley, Gemma L.C.; Forest, Félix; Rogier (2009). "Troublesome tropical mints: re-examining generic limits of Vitex and relations (Lamiaceae) in South East Asia". Taxon 58 (2): 500–510. doi:10.1002/tax.582014. Bibcode: 2009Taxon..58..500B. 
  15. ↑ Scheen, Anne-Cathrine; Albert, Victor A. (2007). "Nomenclatural and taxonomic changes within the Leucas clade (Lamioideae; Lamiaceae)". Systematics and Geography of Plants 77 (2): 229–238. https://www.ingentaconnect.com/content/nbgb/sgp/2007/00000077/00000002/art00004. 
  16. ↑ Scheen, Anne-Cathrine; Albert, Victor A. (2009). "Molecular Phylogenetics of the Leucas Group (Lamioideae; Lamiaceae)". Systematic Botany 34 (1): 173–181. doi:10.1600/036364409787602366. Bibcode: 2009SysBo..34..173S. 
  17. ↑ 17.0 17.1 17.2 Li, Bo; Cantino, Philip D.; Olmstead, Richard G.; Bramley, Gemma L. C.; Xiang, Chun-Lei; Ma, Zhong-Hui; Tan, Yun-Hong; Zhang, Dian-Xiang (2016-10-17). "A large-scale chloroplast phylogeny of the Lamiaceae sheds new light on its subfamilial classification" (in en). Scientific Reports 6 (1). doi:10.1038/srep34343. ISSN 2045-2322. PMID 27748362. PMC 5066227. https://www.nature.com/articles/srep34343. 
  18. ↑ 18.0 18.1 18.2 18.3 Zhao, Fei; Chen, Ya-Ping; Salmaki, Yasaman; Drew, Bryan T.; Wilson, Trevor C.; Scheen, Anne-Cathrine; Celep, Ferhat; Bräuchler, Christian et al. (8 January 2021). "An updated tribal classification of Lamiaceae based on plastome phylogenomics". BMC Biology 19 (1): 2. doi:10.1186/s12915-020-00931-z. PMID 33419433. 
  19. ↑ "Lamiaceae Genera". https://www.mobot.org/mobot/research/apweb/genera/lamiaceaegen.html. 
  20. ↑ Zhong, Jin-Shun; Li, Jie; Li, Lang; Conran, John G.; Hsi-wen, Li (2010). "Phylogeny of Isodon (Schrad. ex Benth.) Spach (Lamiaceae) and Related Genera Inferred from Nuclear Ribosomal ITS, trnL-trnF Region, and rps16 Intron Sequences and Morphology". Systematic Botany 35 (1): 207–219. doi:10.1600/036364410790862614. Bibcode: 2010SysBo..35..207Z. 
  21. ↑ Walker, Jay B.; Sytsma, Kenneth J. (2007). "Staminal Evolution in the Genus Salvia (Lamiaceae): Molecular Phylogenetic Evidence for Multiple Origins of the Staminal Lever". Annals of Botany 100 (2): 375–391. doi:10.1093/aob/mcl176. PMID 16926227. 
  22. ↑ Ryding, P. Olof (2010). "Pericarp structure and phylogeny of tribe Mentheae (Lamiaceae)". Plant Systematics and Evolution 285 (3–4): 165–175. doi:10.1007/s00606-010-0270-9. Bibcode: 2010PSyEv.285..165R. 

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